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A framework for understanding semi-permeable barrier effects on migratory ungulates

1. Impermeable barriers to migration can greatly constrain the set of possible routes and ranges used by migrating animals. For ungulates, however, many forms of development are semi-permeable, and making informed management decisions about their potential impacts to the persistence of migration routes is difficult because our knowledge of how semi-permeable barriers affect migratory behaviour and function is limited. 2. Here, we propose a general framework to advance the understanding of barrier effects on ungulate migration by emphasizing the need to (i) quantify potential barriers in terms that allow behavioural thresholds to be considered, (ii) identify and measure behavioural responses to semi-permeable barriers and (iii) consider the functional attributes of the migratory landscape (e.g. stopovers) and how the benefits of migration might be reduced by behavioural changes. 3. We used global position system (GPS) data collected from two subpopulations of mule deer Odocoileus hemionus to evaluate how different levels of gas development influenced migratory behaviour, including movement rates and stopover use at the individual level, and intensity of use and width of migration route at the population level. We then characterized the functional landscape of migration routes as either stopover habitat or movement corridors and examined how the observed behavioural changes affected the functionality of the migration route in terms of stopover use. 4. We found migratory behaviour to vary with development intensity. Our results suggest that mule deer can migrate through moderate levels of development without any noticeable effects on migratory behaviour. However, in areas with more intensive development, animals often detoured from established routes, increased their rate of movement and reduced stopover use, while the overall use and width of migration routes decreased. 5. Synthesis and applications . In contrast to impermeable barriers that impede animal movement, semi-permeable barriers allow animals to maintain connectivity between their seasonal ranges. Our results identify the mechanisms (e.g. detouring, increased movement rates, reduced stopover use) by which semi-permeable barriers affect the functionality of ungulate migration routes and emphasize that the management of semi-permeable barriers may play a key role in the conservation of migratory ungulate populations.

Journal of Applied Ecology↗

Fish locomotor dynamics quantified using multiple inertial measurement units

Animals exhibit a diversity of locomotor behaviours that vary with body shape, ontogeny, motivation and environment. Documenting these behaviours often relies on recording videos of animals in controlled conditions displaying stereotyped behaviours. By contrast, animal-borne motion sensing data loggers can document diverse behaviours of animals in their natural environment. Traditionally, one data logger is used to infer body dynamics; however, a single location presents an incomplete picture of body movement. Here, we deploy multiple independent inertial measurement units on individual fish to study within-body dynamics and assess the accuracy of this approach compared to traditional video methods. Using a combination of robotic and animal experiments, we accurately measure the timing, direction and magnitude of movement of multiple locations along the body and can recreate accurate three-dimensional midline kinematics without video data. Furthermore, we show that with as few as two data loggers, parameters such as propulsive wavelength can be calculated, even during high-intensity behaviours. Using multiple data loggers has the potential to provide unbiased spatio-temporal estimates of fish kinematics, allowing kinematic studies to be extended to new species, new habitats and a diversity of natural fish locomotor behaviours beyond those traditionally studied in laboratory settings.

Royal Society Open Science↗

Short-term responses to a human-altered landscape do not affect fat dynamics of a migratory ungulate

According to risk-sensitive foraging theory, animals should make foraging decisions that balance nutritional costs and gains to promote fitness. Human disturbance is a form of perceived risk that can prompt avoidance of risky habitat over acquisition of food. Consequently, behavioural responses to perceived risk could induce nutritional costs. Population declines often coincide with increases in human disturbance, which likely is associated with direct and indirect habitat loss. Nevertheless, behavioural and physiological responses to perceived risks associated with human disturbance could be an added nutritional deficit with population-level repercussions. Using GPS-collar data from three populations of migratory mule deer Odocoileus hemionus exposed to a gradient of established industrial energy development on winter ranges where direct and indirect habitat loss were well documented, we evaluated whether exposure and behavioural responses to human disturbance alter changes in nutritional condition (i.e. fat reserves) over winter. Although animals exhibited behaviours indicative of perceived risk of human disturbance, such as increased movement rates and avoidance of infrastructure, exposure and behavioural responses to human disturbance had little to no measurable effect on fat loss over a winter. Instead, catabolism of fat reserves occurred primarily as a function of the amount of fat animals had entering winter, suggesting that, in the short term, animals were able to mitigate the energetic costs of perceived risk of human disturbance over winter. Animals, however, did not appear to overcome persistent food limitations, and animals with less food availability lost more fat over winter. Our findings heed caution in using short-term behavioural and physiological responses to inform long-term nutritional consequences of human disturbance. Although animals appear to mitigate the energetic costs of perceived risk, food limitations exacerbated by broader-scale avoidance of food near human disturbance may be the primary pathway causing the frequently observed population declines following human disturbance to pristine landscapes.

Wyoming↗

Long-term video surveillance and automated analyses reveal arousal patterns in groups of hibernating bats

Understanding natural behaviours is essential to determining how animals deal with new threats (e.g. emerging diseases). However, natural behaviours of animals with cryptic lifestyles, like hibernating bats, are often poorly characterized. White-nose syndrome (WNS) is an unprecedented disease threatening multiple species of hibernating bats, and pathogen-induced changes to host behaviour may contribute to mortality. To better understand the behaviours of hibernating bats and how they might relate to WNS, we developed new ways of studying hibernation across entire seasons. We used thermal-imaging video surveillance cameras to observe little brown bats ( Myotis lucifugus ) and Indiana bats ( M. sodalis ) in two caves over multiple winters. We developed new, sharable software to test for autocorrelation and periodicity of arousal signals in recorded video. We processed 740 days (17,760 hr) of video at a rate of >1,000 hr of video imagery in less than 1 hr using a desktop computer with sufficient resolution to detect increases in arousals during midwinter in both species and clear signals of daily arousal periodicity in infected M. sodalis . Our unexpected finding of periodic synchronous group arousals in hibernating bats demonstrate the potential for video methods and suggest some bats may have innate behavioural strategies for coping with WNS. Surveillance video and accessible analysis software make it now practical to investigate long-term behaviours of hibernating bats and other hard-to-study animals.

Methods in Ecology and Evolution↗

Mechanisms and function of school formation in subyearling American shad (Alosa sapidissima)

Summary The behavioural repertoire of subyearling American shad ( Alosa sapidissima ) was studied in the laboratory to examine pathways leading to Aggregation and School formation at various group sizes (3–100 fish per 500-liter tank). Three hypotheses were tested: (1) schooling is the dominant behavioural activity of subyearling American shad, (2) development and maintenance of cohesive social behaviour is affected b group size, and (3) survival is a function of group size. Seventeen discrete behaviour patterns were observed. In relation to group size, the amount of time spent Schooling varied directly whereas Following and Aggregating varied inversely. The number of bouts of Parallel orient, Parallel swim, and Pivot also varied inversely with group size. A conceptual model was developed and tested to show how normal schooling behaviour is achieved in small or fragmented groups through intermediary modal action patterns. Sequence analysis showed behavioural diversity to diminish with group size, thereby increasing group cohesion and uniformity of response to stimuli. Probability of survival, determined from weekly mortality in experimental tanks, increased significantly with group size (0.87–0.98 for 3–100 fish). It is hypothesized that survival improves with an increase in the proportion of time spent Schooling, which may increase feeding efficiency, reduce stress, or reduce metabolic expenditures.

Journal of Applied Ichthyology↗

Use of mammal manure by nesting burrowing owls: a test of four functional hypotheses

Animals have evolved an impressive array of behavioural traits to avoid depredation. Olfactory camouflage of conspicuous odours is a strategy to avoid depredation that has been implicated only in a few species of birds. Burrowing owls, Athene cunicularia, routinely collect dried manure from mammals and scatter it in their nest chamber, in the tunnel leading to their nest and at the entrance to their nesting burrow. This unusual behaviour was thought to reduce nest depredation by concealing the scent of adults and juveniles, but a recent study suggests that manure functions to attract arthropod prey. However, burrowing owls routinely scatter other materials in the same way that they scatter manure, and this fact seems to be at odds with both of these hypotheses. Thus, we examined the function of this behaviour by testing four alternative hypotheses. We found no support for the widely cited olfactory-camouflage hypothesis (manure did not lower the probability of depredation), or for the mate-attraction hypothesis (males collected manure after, not before, pair formation). Predictions of the burrow-occupied hypothesis (manure indicates occupancy to conspecifics and thereby reduces agonistic interactions) were supported, but results were not statistically significant. Our results also supported several predictions of the prey-attraction hypothesis. Pitfall traps at sampling sites with manure collected more arthropod biomass (of taxa common in the diet of burrowing owls) than pitfall traps at sampling sites without manure. Scattering behaviour of burrowing owls appears to function to attract arthropod prey, but may also signal occupancy of a burrow to conspecifics. ?? 2006 The Association for the Study of Animal Behaviour.

Animal Behaviour↗

Changes in vocal repertoire of the Hawaiian crow, Corvus hawaiiensis,from past wild to current captive populations

For most avian species, social behaviou r is critically important for survival and reproductive success. Many social behaviours in birds are culturally transmitted, and as bird populations decline across the globe, important elements of these behaviours may be lost. The Hawaiian crow or 'alalā, Corvus hawaiiensis , is a socially complex avian species that is currently extinct in the wild. As in other oscine passerines, vocalizations in the 'alalā may be culturally transmitted. We compared the vocal repertoire of three of the last four wild 'alalā pairs from the early 1990s to three current captive pairs on the Island of Hawai'i to determine how acoustic behaviour has been affected by changes in their social and physical environment. Over 18 h of recordings from wild breeding pairs were analysed and compared with 44 h from captive breeding pairs. Calls were placed into five putative behavioural categories: (1) alarm, (2) territorial broadcast, (3) aggression, (4) submission and (5) courtship. There was little difference in the overall number and diversity of call types among wild versus captive birds. However, the repertoire was significantly different. Territorial broadcast calls, common components of the wild repertoire, were absent from the captive repertoire. In addition, wild birds had twice the number of alarm calls and had a higher call rate than captive birds. Our results show how socially learned behaviours may change over relatively short periods for an entire species. Understanding how the vocal repertoire and the functional context of vocalizations change may provide useful information for ongoing efforts to reintroduce the 'alalā into the wild.

Hawai'i↗

Look what the cat dragged in: do parasites contribute to human cultural diversity?

If human culture emerges from the modal personality of a population, can global variation in parasitism that affects personality lead to cultural diversity among nations? The answer could help explain why people seem to vary so much from one land to another. Thomas et al. (2005) review how parasites manipulate behaviour, including human behaviour. To quote them, “The rabies virus lives in the brain, affording the virus ample opportunity to directly affect host behaviour. Rabid animals do show changes in behaviour, including increased aggression and biting.” Rabies affects a wide range of mammals and the aggressive biting associated with furious rabies appears to increase transmission. The personality transformation of infected humans can be horrifying, transforming loved ones into thrashing, baying beasts. Not coincidentally, in Europe, past periods of rabies outbreaks correspond to increases in werewolf trials. Although rabies can have a dramatic effect, the present rarity of human rabies cases and the availability of a vaccine, means that the behavioural effects of rabies are primarily an illustrative curiosity.

Behavioural Processes↗

Performance comparisons for artificially propagated and wild Pacific lamprey juveniles and larvae

Artificially propagated Pacific lamprey ( Entosphenus tridentatus ) are produced for restoration and for use in dam passage studies to reduce the demand for wild fish. Such uses require that animals are representative of their wild counterparts. Previous work indicated that this is true for Pacific lamprey larvae and juveniles reared in the hatchery with respect to the length of sustained swimming. However, more subtle differences in behaviour and performance that lamprey need to survive have not been assessed. In this study, artificially propagated and wild fish were compared in laboratory tests under no-flow conditions to examine light avoidance, burrowing speed, burst swim speed, volitional routine swim speed and time to come to rest. Most larvae burrowed in less than a minute, and we found highly significant differences ( p < 0.001) between artificially propagated and wild larvae burrowing times, a critical escape behaviour. This could have implications for studies of larval entrainment at irrigation diversion canals or in turbine boils at dams. Interestingly, all of the wild juveniles tested came to rest quickly after introduction to the chamber (1.5 min), while artificially propagated lamprey swam robotically near the surface and 48% did not come to rest in the first 10 min (median time to rest = 9.5 min). In contrast, wild juveniles quickly (median = 1.47 min) sought areas near the tank bottom and attached strongly with their oral disc. Such behavioural differences could have important survival consequences for artificially propagated lamprey as they approach turbine intakes, bypass screens and irrigation diversion headgates. This study highlights the need to conduct behavioural assays that examine subtleties of fish behaviour that can be missed with traditional swim tunnel comparisons.

Aquaculture, Fish and Fisheries↗

Conflict of interest between a nematode and a trematode in an amphipod host: Test of the "sabotage" hypothesis

Microphallus papillorobustus is a manipulative trematode that induces strong behavioural alterations in the gamaridean amphipod Gammarus insensibilis , making the amphipod more vulnerable to predation by aquatic birds (definitive hosts). Conversely, the sympatric nematode Gammarinema gammari uses Gammarus insensibilis as a habitat and a source of nutrition. We investigated the conflict of interest between these two parasite species by studying the consequences of mixed infection on amphipod behaviour associated with the trematode. In the field, some amphipods infected by the trematode did not display the altered behaviour. These normal amphipods also had more nematodes, suggesting that the nematode overpowered the manipulation of the trematode, a strategy that would prolong the nematode's life. We hypothesize that sabotage of the trematode by the nematode would be an adaptive strategy for the nematode consistent with recent speculation about co-operation and conflict in manipulative parasites. A behavioural test conducted in the laboratory from naturally infected amphipods yielded the same result. However, exposing amphipods to nematodes did not negate or decrease the manipulation exerted by the trematode. Similarly, experimental elimination of nematodes from amphipods did not permit trematodes to manipulate behaviour. These experimental data do not support the hypothesis that the negative association between nematodes and manipulation by the trematode is a result of the "sabotage" hypothesis.

Behavioral Ecology and Sociobiology↗

Snowshoe hares display limited phenotypic plasticity to mismatch in seasonal camouflage

As duration of snow cover decreases owing to climate change, species undergoing seasonal colour moults can become colour mismatched with their background. The immediate adaptive solution to this mismatch is phenotypic plasticity, either in phenology of seasonal colour moults or in behaviours that reduce mismatch or its consequences. We observed nearly 200 snowshoe hares across a wide range of snow conditions and two study sites in Montana, USA, and found minimal plasticity in response to mismatch between coat colour and background. We found that moult phenology varied between study sites, likely due to differences in photoperiod and climate, but was largely fixed within study sites with only minimal plasticity to snow conditions during the spring white-to-brown moult. We also found no evidence that hares modify their behaviour in response to colour mismatch. Hiding and fleeing behaviours and resting spot preference of hares were more affected by variables related to season, site and concealment by vegetation, than by colour mismatch. We conclude that plasticity in moult phenology and behaviours in snowshoe hares is insufficient for adaptation to camouflage mismatch, suggesting that any future adaptation to climate change will require natural selection on moult phenology or behaviour.

Proceedings of the Royal Society B: Biological Sci↗

On the interface between cultural transmission, phenotypic diversity, demography and the conservation of migratory ungulates

Recent evidence indicates that green-wave surfing behaviour in ungulates and the migrations that stem from this behaviour are socially learned, culturally transmitted across generations and become more efficient via cumulative cultural evolution. But given a lack of corroborative evidence, whether ungulate migration is a cultural phenomenon remains a hypothesis deserving of further testing. In this opinion piece, we summarize the role memory and social learning play in the green-wave surfing that underlies ungulate migration, and when combined with the natural history of ungulates, we argue that the most likely mechanism for maintenance of ungulate migration is animal culture. We further our argument by providing a synopsis of processes that promote diversification of migratory behaviour and link these processes to their emergent ecological patterns, which are common in nature but have not historically been considered as potential cultural phenomena. The notion that diverse portfolios of migratory behaviour may buffer populations from environmental change emerges from this synthesis but requires empirical testing. Finally, we contend that, because the migratory behaviour of ungulates stems largely from cultural transmission as opposed to a genetic programme, the diversity of observed migratory strategies represents ‘culturally significant units’ deserving of the same conservation effort afforded to evolutionarily significant units.

Wyoming↗

Threading the needle: How humans influence predator–prey spatiotemporal interactions in a multiple‐predator system

Perceived predation risk and the resulting antipredator behaviour varies across space, time and predator identity. Communities with multiple predators that interact and differ in their use of space, time of activity and hunting mode create a complex landscape for prey to avoid predation. Anthropogenic presence and disturbance have the potential to shift interactions among predators and prey and the where and when encounters occur. We examined how white-tailed deer Odocoileus virginianus fawn spatiotemporal antipredator behaviour differed along an anthropogenic disturbance gradient that had black bears Ursus americanus , coyotes Canis latrans , bobcats Lynx rufus and humans present. We quantified (a) spatial co-occurrence in species distributions, (b) temporal overlap across the diel cycle and (c) spatiotemporal associations between humans, bears, coyotes, bobcats, adult male deer and fawns. We also examined how deer vigilance behaviour changed across the anthropogenic disturbance gradient and survey duration. Anthropogenic disturbance influenced spatiotemporal co-occurrence across multiple scales, often increasing spatiotemporal overlap among species. In general, species’ spatial co-occurrence was neutral or positive in anthropogenically disturbed environments. Bears and fawns, coyotes and adult male deer, and bobcats and fawns all had higher temporal overlap in the agriculture-development matrix sites. In addition, factors that influenced deer vigilance (e.g. distance to forest edge and predator relative abundance) in the agriculture-development matrix sites did not in the forest matrix site. By taking into account the different antipredator behaviours that can be detected and the different scales these behaviours might occur, we were able to gain a more comprehensive picture of how humans reduce available niche space for wildlife, creating the neutral and positive spatiotemporal associations between species that studies have been seeing in more disturbed areas.

Pennsylvania↗

Behavioral state-dependent habitat selection and implications for animal translocations

Post-release monitoring of translocated animals is often used to inform future translocation protocols. Quantifying habitat selection of translocated individuals may help identify features that characterize good settlement habitat and thus inform the choice of future release sites. However, translocated animals often undergo post-release behavioural modification, and their habitat selection may vary depending on the underlying behavioural state. To investigate this, we analysed behavioural state-dependent habitat selection in female greater sage-grouse Centrocercus urophasianus translocated from Wyoming to North Dakota, USA, using Hidden Markov Models combined with Integrated Step Selection Analysis. We segmented individual trajectories into behavioural phases corresponding to an exploratory state, characterized by broad and directed movements, and a restricted state, characterized by short and tortuous movements. Then, we quantified habitat selection in each state while accounting for seasonality and individual reproductive status. While in the exploratory state, sage-grouse exhibited natal habitat preference induction by selecting for high sagebrush cover, which is typical of their natal area in Wyoming but not of the release area in North Dakota. In the restricted state, sage-grouse selected for gentle topography and also adjusted their habitat selection to constraints imposed by seasonality and reproductive needs by selecting for high herbaceous cover during brood rearing. Synthesis and applications . Habitat selection of translocated sage-grouse differed between the post-release exploration and the settlement phase. Features selected after settling, not during exploration, are likely indicative of suitable settlement habitat. Our results suggest that areas characterized by gentle topography and high herbaceous cover are well-suited as release sites for sage-grouse translocated during the summer, especially brood-rearing females, and that sagebrush cover may not be a critical factor in determining the appropriateness of release sites for sage-grouse in North Dakota. Our findings highlight the need to consider behaviour when using habitat selection estimates to inform the choice of future release sites.

North Dakota, Wyoming↗

Factors influencing Cinnamon Teal nest attendance patterns

Patterns of nest attendance in birds result from complex behaviours and influence the success of reproductive events. Incubation behaviours vary based on individual body condition, energy requirements and environmental factors. We assessed nest attendance patterns in Cinnamon Teal Spatula cyanoptera breeding in the San Luis Valley of Colorado in 2016–2017 using trail and video cameras to observe behaviours throughout incubation. We evaluated the effect of temporal, life-history and environmental covariates on the frequency and duration of incubation recesses as well as the incubation constancy. There was considerable model uncertainty among the models used to evaluate recess frequency. Recess duration varied according to the interaction between nest age and a quadratic effect of time of day, with hens on older nests taking longer recesses in the afternoon and hens on nests earlier in incubation taking longer recesses in the morning and evening. Incubation constancy decreased with higher ambient temperatures in the study area. This study provides evidence that Cinnamon Teal modify their behaviour during incubation according to the age of the nest and the time of day. These results improve our knowledge of Cinnamon Teal breeding ecology and shed light on the behaviours that fast-lived species may use to cope with environmental factors during nesting.

Colorado↗

Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasianus , populations

S age grouse, Centrocercus urophasianus , in an isolated montane basin near Gunnison, Colorado differ in several morphological and behavioural traits from conspecifics studied in other areas of the species' range. Both sexes in Gunnison are smaller than sage grouse elsewhere, and males possess differences in feather morphology as well. The mating behaviour of male sage grouse in three populations was examined to determine whether male strut displays of Gunnison sage grouse were behaviourally distinct. Behavioural analyses revealed Gunnison males perform strut displays at a slower rate than males in the two other sage grouse populations sampled. In addition, Gunnison males' strut displays contain unique visual and acoustical aspects. The most distinguishing attributes of Gunnison sage grouse were male secondary sexual characteristics including traits that correlate with mating success in other populations. Thus, phenotypic differences observed in the Gunnison population represent a divergence in expression of traits that are likely to be influenced by sexual selection. Recent models of speciation suggest that species characterized by intense sexual selection, such as those with lek mating systems, have the potential for rapid inter-populational divergence in male traits and female preferences leading to speciation.

Colorado↗

A new approach to the study of relationship quality in dolphins: framework and preliminary results

Proximity and synchronous behaviours from surface observations have been used to measure association patterns within and between dolphin dyads. To facilitate an investigation of relationship quality in dolphins, we applied a method used for chimpanzees and ravens that examined three main components to describe relationships: value, security, and compatibility. Using pilot data from a long-term study of two study populations for this preliminary assessment, these three components were extracted from PCA analysis of eight behavioural variables with more than 80% variance accounted for in both study groups. Only pair swim position differed between groups. Although value, security, and compatibility are abstract terms, each is based on behaviours identified as important in dolphin social life, at least for these two populations. Examining relationship quality in dolphins with a method used to illustrate dyadic differences for chimpanzees and ravens allows for a quantitative, comparative assessment of sociality across disparate taxa. Although these three species are diverse in their anatomies and in their social habitats (e.g., aquatic, terrestrial, aerial), they may well share the basic societal building blocks in the factors affecting how relationships are formed. We discuss how an examination of these behavioural variables facilitates understanding relationship quality in dolphins, as well as how dolphin relationships fit into the context of social animals’ society.

Bimini Islands, Great Bahama Bank, Roatan Island↗

Quantifying the behavioral response of spawning chum salmon to elevated discharges from Bonneville Dam, Columbia River, USA

Chum salmon Oncorhynchus keta that spawn in main-stem habitats below Bonneville Dam on the Columbia River, USA, are periodically subjected to elevated discharges that may alter spawning behaviour. We investigated behavioural responses of spawning chum salmon to increased water velocities associated with experimental increases in tailwater elevation using acoustic telemetry and a dual-frequency identification sonar. Chum salmon primarily remained near their redds at base tailwater elevations (3.5 m above mean sea level), but displayed different movement and behavioural responses as elevations were increased to either 4.1 or 4.7m for 8-h periods. When velocities remained suitable (<0.8m s -1 ) during elevated-tailwater tests, female chum salmon remained near their redds but exhibited reduced digging activity as water velocities increased. However, when velocities exceeded 0.8m s -1 , the females that remained on their redds exhibited increased swimming activity and digging virtually ceased. Female and male chum salmon that left their redds when velocities became unsuitable moved mean distances ranging from 32 to 58 m to occupy suitable velocities, but returned to their redds after tailwaters returned to base levels. Spawning events (i.e. egg deposition) were observed for five of nine pairs of chum salmon following tests indicating any disruptions to normal behaviour caused by elevated tailwaters were likely temporary. We believe a chum salmon's decision to either remain on, or leave, its redd during periods of unsuitably high water velocities reflects time invested in the redd and the associated energetic costs it is willing to incur. ?? 2009 John Wiley & Sons, Ltd.

River Research and Applications↗